Oviraptorosaurs ("egg thief lizards") are a group of feathered maniraptoran dinosaurs from the Cretaceous Period of what are now Asia and North America. They are distinct for their characteristically short, beaked, parrot-like skulls, with or without bony crests atop the head. They ranged in size from Caudipteryx, which was the size of a turkey, to the 8 metre long, 1.4 ton Gigantoraptor. The group (along with all maniraptoran dinosaurs) is close to the ancestry of birds. Analyses like those of Maryanska et al (2002) and Osmólska et al. (2004) suggest that they may represent primitive flightless birds. The most complete oviraptorosaur specimens have been found in Asia. The North American oviraptorosaur record is sparse.
The earliest and most basal ("primitive") known oviraptorosaurs are Ningyuansaurus wangi, Protarchaeopteryx robusta and Incisivosaurus gauthieri, both from the lower Yixian Formation of China, dating to about 125 million years ago during the Aptian age of the early Cretaceous period. A tiny neck vertebra reported from the Wadhurst Clay Formation of England shares some features in common with oviraptorosaurs, and may represent an earlier occurrence of this group (at about 140 million years ago).
|Replica of a Caudipteryx zoui skeleton|
Oviraptorosaurians have shortened rostrums, massive, beaklike mandibles, and long parietal bones. With the exception of the 8-meter long Gigantoraptor, they are generally medium-sized and rarely exceeded 2 meters in length. The most primitive members have four pairs of teeth in the premaxillae, such as in Caudipteryx and in Incisivosaurus they are enlarged and form bizarrely prominent bucktoothed incisors. The more advanced members have no teeth in the jaws.
Pneumatization is extensive in the skulls and vertebrae of the more advanced members. Oviraptorosauria have thick, U - shaped furculae and a large sternal plates that are wider (together) than they are long, unlike in birds and dromaeosaurs. The arms are around half the length of the legs and over half the length of the presacral vertebral column. The hands are long, and tridactyl, with a reduced third finger in Caudipteryx and Ajancingenia. There are between 5 and 8 sacral vertebrae. The pubis is vertical or subvertical, with a concave anterior edge. The tibia is 15%-25% longer than the femur. The tail is short, with the number of vertebrae reduced to 24 or so, and proximally very thick, with broad transverse processes. The ischium retains the primitive character of a prominent, triangular obturator process and lack the proximodorsal process that is found in birds. In advanced oviraptorosaurians, the ischium is curved posteriorly. The pectoral girdle is also primitive; the scapula is a broad blade that is distally expanded, it lies on the lateral aspect of the thorax at an angle to the vertebral column, and the coracoid has the primitive coelurosaur shape with a proximal supracoracoidal nerve foramen and a moderate biceps tubercle.
Oviraptorosaurians are different from most other maniraptorans in the form of their skulls. They have shortened snouts, beak-like jaws with few or no teeth, and a large opening in the lower jaw bone. Some have bony crests atop the skull. The most primitive members have a few teeth in the front of the mouth; in Incisivosaurus, they are enlarged and form bizarrely prominent "bucktoothed" incisors. The arms and hands are generally long (though very reduced in some advanced species) and the shoulder girdle is large and massive, with flexed coracoid bones and prominent attachments for strong arm muscles.
Their tails are very short compared to other maniraptorans. In Nomingia and Similicaudipteryx, the tail ends in four fused vertebrae which Osmólska, He, and others have referred to as a "pygostyle", but which Witmer found was anatomically different and evolved separately from the pygostyle of birds (a bone which serves as the attachment point for a fan of tail feathers).
Evidence for feathered oviraptorosaurs exists in several forms. Most directly, four species of primitive oviraptorosaurs (in the genera Caudipteryx, Protarchaeopteryx, and Similicaudipteryx) have been found with impressions of well developed feathers, most notably on the wings and tail, suggesting that they functioned at least partially for display. Secondly, at least four oviraptorosaur genera (Nomingia, Similicaudipteryx, Citipati, and Conchoraptor) preserved tails ending in something like a pygostyle, a bony structure at the end of the tail that, in modern birds, is used to support a fan of feathers. Similarly, quill knobs (anchor points for wing feathers on the ulna) have been reported in the oviraptorosaurian species Avimimus portentosus. Additionally, a number of oviraptorid specimens have famously been discovered in a nesting position similar to that of modern birds. The arms of these specimens are positioned in such a way that they could perfectly cover their eggs if they had small wings and a substantial covering of feathers.
The eating habits of these animals are not fully known: they have been suggested to have been either carnivorous, herbivorous, mollusk-eating or egg-eating (the evidence that originally supported the latter is no longer considered valid); these options are not necessarily incompatible.
Some ate small vertebrates. Evidence for this comes from a lizard skeleton preserved in the body cavity of Oviraptor and two baby Troodontid skulls found in a Citipati nest. Evidence in favor of a herbivorous diet includes the presence of gastroliths preserved with Caudipteryx. There are also arguments for the inclusion of mollusks in their diet.
Originally these animals were thought to be egg raiders, based on a Mongolian find showing Oviraptor on top of a nest. Recent studies have shown that the animal was actually on top of its own nest.
Several oviraptorosaurian nests are known, with several oviraptorid specimens preserved in a brooding position over large clutches of up to a dozen or more eggs. The eggs are usually arranged in pairs, and forming a circular pattern within the nest. One oviraptorosaurian specimen from China has been found with two unlaid eggs within the pelvic canal. This suggests that, unlike modern crocodilians, oviraptorosaurs did not produce and lay many eggs at the same time. Rather, the eggs were produced within the reproductive organs in pairs, and laid two at a time, with the mother positioned in the center of the nest and rotating in a circle as each pair was laid. This behavior is supported by the fact that the eggs were shaped like highly elongated ovals, with the more pointed end pointing backward from the cloaca, and also oriented toward the center of the nest. Geochemical analysis also revealed that oviraptorosaurs incubated their eggs in the 35–40 °C (95–104 °F) range, as many modern bird species do today, based on the oxygen isotope ratios in the bones of the fossil embryos of various species during development.
The presence of two shelled eggs within the birth canal shows that oviraptorosaurs were intermediate between the reproductive biology of crocodilians and modern birds. Like crocodilians, they had two oviducts. However, crocodilians produce multiple shelled eggs per oviduct at a time, whereas oviraptorosaurs, like birds, produced only one egg per oviduct at a time.
Oviraptorosaurs, like deinonychosaurs, are so bird-like that several scientists consider them to be true birds, more advanced than Archaeopteryx. Gregory S. Paul has written extensively on this possibility, and Teresa Maryańska and colleagues published a technical paper detailing this idea in 2002. Michael Benton, in his widely respected text Vertebrate Paleontology, also included oviraptorosaurs as an order within the class Aves. However, a number of researchers have disagreed with this classification, retaining oviraptorosaurs as non-avialan maniraptorans slightly more primitive than the deinonychosaurs.
The internal classification of the oviraptorosaurs has also been controversial. Most studies divide them into two primary sub-groups, the Caenagnathidae and the Oviraptoridae. The Oviraptoridae is further divided into the small, short-armed, and crestless subfamily Ingeniinae, and the larger, crested, long-armed Oviraptorinae. However, some phylogenetic studies have suggested that many traditional members of the Caenagnathidae were more closely related to the crested oviraptorids.
The 2007 cladistic analysis of Turner and colleagues recovered the Oviraptorosauria as a maniraptoran clade (natural grouping) of maniraptorans more primitive than true birds. They found that the oviraptorosaurs are the sister group to the Therizinosauria and that the two, together, are more basal than any member of Paraves. However, a more recent study by Zanno and colleagues challenged that finding, showing therizinosaurs to be more primitive and not closely related to oviraptorosaurs.
The following cladogram was found by an analysis published with the description of the caenagnathid Anzu.
Apatoraptor ("Apatè robber") is a genus of caenagnathid dinosaur which contains a single species, A. pennatus. The only known specimen was discovered in the Campanian-age Horseshoe Canyon Formation of Alberta.Caenagnathasia
Caenagnathasia ('recent jaw from Asia') is a small caenagnathid oviraptorosaurian theropod from the Late Cretaceous of Uzbekistan.Caenagnathidae
Caenagnathidae is a family of bird-like maniraptoran theropod dinosaurs from the Late Cretaceous of North America and Asia. They are a member of the Oviraptorosauria, and close relatives of the Oviraptoridae. Like other oviraptorosaurs, caenagnathids had specialized beaks, long necks, and short tails, and would have been covered in feathers. The relationships of caenagnathids were long a puzzle. The family was originally named by Charles Hazelius Sternberg in 1940 as a family of flightless birds. The discovery of skeletons of the related oviraptorids revealed that they were in fact non-avian theropods, and the discovery of more complete caenagnathid remains revealed that Chirostenotes pergracilis, originally named on the basis of a pair of hands, and "Ornithomimus" elegans, named from a foot, were caenagnathids as well.Caudipteryx
Caudipteryx (which means "tail feather") is a genus of peacock-sized theropod dinosaurs that lived in the Aptian age of the early Cretaceous Period (about 124.6 million years ago). They were feathered and remarkably birdlike in their overall appearance.
Two species have been described; C. zoui (the type species), in 1998, and C. dongi, in 2000.Caudipteryx fossils were first discovered in the Yixian Formation of the Sihetun area of Liaoning Province, northeastern China in 1997.Coelurosauria
Coelurosauria (; from Greek, meaning "hollow tailed lizards") is the clade containing all theropod dinosaurs more closely related to birds than to carnosaurs.
Coelurosauria is a subgroup of theropod dinosaurs that includes compsognathids, tyrannosaurs, ornithomimosaurs, and maniraptorans; Maniraptora includes birds, the only dinosaur group alive today.Most feathered dinosaurs discovered so far have been coelurosaurs. Philip J. Currie considers it probable that all coelurosaurs were feathered. In the past, Coelurosauria was used to refer to all small theropods, but this classification has since been abolished.Elmisaurus
Elmisaurus is an extinct genus of dinosaur from the Late Cretaceous. It was a theropod belonging to the Oviraptorosauria. Its fossils have been found in Mongolia. It is known from foot and hand bones.Epidexipteryx
Epidexipteryx is a genus of small paravian dinosaurs, known from one fossil specimen in the collection of the Institute of Vertebrate Paleontology and Paleoanthropology in Beijing. Epidexipteryx represents the earliest known example of ornamental feathers in the fossil record. The type specimen is catalog number IVPP V 15471. It has been reported to be a maniraptoran dinosaur from the Middle Jurassic or Upper Jurassic age Daohugou Beds of Inner Mongolia, China (about 160 or 168 mya).The specific name, Epidexipteryx hui ("Hu's display feather"), and its Chinese name Hushi Yaolong ("Hu Yaoming's dragon") were coined in memory of paleomammologist Hu Yaoming.Hagryphus
Hagryphus ("Ha's griffin"), is an oviraptorosaurian theropod dinosaur from the Upper Cretaceous Period of what is now Utah.Heyuannia
Heyuannia ("from Heyuan") is a genus of oviraptorid dinosaur that lived during the Late Cretaceous Period in China. It was the first oviraptorid found in that country; most others were found in neighbouring Mongolia. Two species are known: H. huangi, named by Lü Junchang in 2002; and H. yanshini, originally named as a separate genus Ingenia by Rinchen Barsbold in 1981 (and renamed to Ajancingenia in 2013 due to the preoccupation of Ingenia).Incisivosaurus
Incisivosaurus ("incisor lizard") is a genus of small, probably herbivorous theropod dinosaur from the early Cretaceous Period of what is now the People's Republic of China. The first specimen to be described (by Xu et al. in 2002), IVPP V13326, is a skull that was collected from the lowermost levels (the fluvial Lujiatun beds) of the Yixian Formation (dating to the Barremian stage about 126 million years ago in the Sihetun area, near Beipiao City, in western Liaoning Province. The most significant, and highly unusual, characteristic of this dinosaur is its apparent adaptation to an herbivorous or omnivorous lifestyle. It was named for its prominent, rodent-like front teeth, which show wear patterns commonly found in plant-eating dinosaurs. The specific name gauthieri honors Dr. Jacques Gauthier, a pioneer of the phylogenetic method of classification.Kakuru
Kakuru is a genus of theropod dinosaur from the early Cretaceous Period.
Kakuru is known primarily from a single fossilised tibia, which had been fossilised through a rare process in which the bone through hydration turned to opal. The bone was dug up at the opal fields of Andamooka, South Australia. The opalised tibia was exhibited by a gem shop in 1973 and by chance brought to the attention of paleontologist Neville Pledge. The owner at the time, a certain A. Fleming, allowed pictures and two casts to be made but eventually the specimen was sold at an auction to an anonymous buyer. It was presumed lost to science. In 2004, however, the South Australian Museum succeeded in procuring the fossil for $22,000.
Kakuru was formally named in 1980 by Pledge and Ralph Molnar. The type species is Kakuru kujani. The generic name is that of a Rainbow Serpent of Australian Aboriginal mythology. The specific name is that of the local aboriginal tribe, the Kujani or Guyani.
One of the casts is the plastoholotype, SAM P17926. The specimen was discovered in the marine Marree Formation dating from the Aptian. Apart from the tibia, the find included some small probable fibula fragments. Later a foot digit was referred that might have come from the same species, specimen SAM P18010, but the assignment is dubious. The tibia is broken into about ten larger pieces and roughly 33 centimetres long. It is very slender in build and shows the impression of the ascending process of the astragalus, an ankle bone itself lost. The process seems to have been very long and narrow.
Kakuru is believed to have been carnivorous, was bipedal and about two to three meters in length. This small dinosaur seems to have had long, slender legs.
Due to the paucity of the remains it has been difficult to establish the phylogenetic position of Kakuru. Molnar and Pledge gave no more precise determination than a Theropoda incertae sedis. The tibia provides two indications: its gracile form and the tall height and narrow width of the astragalar process. Both seems to point to the Coelurosauria, perhaps the Oviraptorosauria. However, in 2005 Oliver Rauhut pointed out that the Abelisauroidea also have a high ascending process of the astragalus.Maniraptora
Maniraptora is a clade of coelurosaurian dinosaurs that includes the birds and the non-avian dinosaurs that were more closely related to them than to Ornithomimus velox. It contains the major subgroups Avialae, Deinonychosauria, Oviraptorosauria and Therizinosauria. Ornitholestes and the Alvarezsauroidea are also often included. Together with the next closest sister group, the Ornithomimosauria, Maniraptora comprises the more inclusive clade Maniraptoriformes. Maniraptorans first appear in the fossil record during the Jurassic Period (see Eshanosaurus), and are regarded as surviving today as living birds.Nankangia
Nankangia is an extinct genus of caenagnathoid oviraptorosaurian dinosaur known from the Late Cretaceous Nanxiong Formation of Nankang County, Ganzhou City of Jiangxi Province, southeastern China. It contains a single species, Nankangia jiangxiensis. N. jiangxiensis coexisted with at least four other caenagnathoids, including an unnamed oviraptorid, Banji long, Ganzhousaurus nankangensis and Jiangxisaurus ganzhouensis. The relatively short dentary and non-downturned mandibular symphysis of Nankangia suggest that it may have been more herbivorous than carnivorous.Nanxiong Formation
The Nanxiong Formation is a Late Cretaceous geologic formation in Guangdong Province. Dinosaur remains are among the fossils that have been recovered from the formation. It consists of red sandstone and clay sediments, with fauna which is more similar to that of the Nemegt Formation in the recent years.Nomingia
Nomingia is a genus of oviraptorid theropod dinosaur hailing from the Late Cretaceous Bugin Tsav Beds of Mongolia.Pennaraptora
Pennaraptora (Latin penna "bird feather" + raptor "thief", from rapere "snatch"; a feathered bird-like predator) is a clade defined as the most recent common ancestor of Oviraptor philoceratops, Deinonychus antirrhopus, and Passer domesticus (the house sparrow), and all descendants thereof, by Foth et al., 2014. The earliest known definitive member of this clade is Anchiornis, from the late Jurassic period of China, about 160 million years ago.
The clade "Aviremigia" was conditionally proposed along with several other apomorphy-based clades relating to birds by Jacques Gauthier and Kevin de Queiroz in a 2001 paper. Their proposed definition for the group was "the clade stemming from the first panavian with ... remiges and rectrices, that is, enlarged, stiff-shafted, closed-vaned (= barbules bearing hooked distal pennulae), pennaceous feathers arising from the distal forelimbs and tail".Protarchaeopteryx
Protarchaeopteryx (meaning "before Archaeopteryx") is a genus of turkey-sized feathered theropod dinosaur from China. Known from the Jianshangou bed of the Yixian Formation, it lived during the early Aptian age of the Early Cretaceous, approximately 124.6 million years ago. It was probably an herbivore or omnivore, although its hands were very similar to those of small carnivorous dinosaurs. It appears to be one of the most basal members of the Oviraptorosauria, closely related to or synonymous with Incisivosaurus.Rinchenia
Rinchenia is a genus of Mongolian oviraptorid dinosaur from the late Cretaceous Period. The type (and only known) species, Rinchenia mongoliensis, was originally classified as a species within the genus Oviraptor (named Oviraptor mongoliensis by Rinchen Barsbold in 1986), but a re-examination by Barsbold in 1997 found differences significant enough to warrant a separate genus. The name Rinchenia was coined for this new genus by Barsbold in 1997, though he did not describe it in detail, and the name remained a nomen nudum until used by Osmólska et al. in 2004.Rinchenia is known from a single specimen (GI 100/32A) consisting of a complete skull and lower jaw, partial vertebral column, partial forelimbs and shoulder girdle, partial hind limbs and pelvis, and a furcula ("wishbone"). While Rinchenia was about the same size as Oviraptor (about 1.5 meters, or 5 ft long), several features of its skeleton, especially in the skull, show it to be distinct. Its skeleton was more lightly built and less robust than that of Oviraptor, and while the crest of Oviraptor is indistinct because of poor fossil preservation, Rinchenia had a well-preserved, highly developed, dome-like casque which incorporated many bones in the skull that are free of the crest in Oviraptor.Timeline of oviraptorosaur research
This timeline of oviraptorosaur research is a chronological listing of events in the history of paleontology focused on the oviraptorosaurs, a group of beaked, bird-like theropod dinosaurs. The early history of oviraptorosaur paleontology is characterized by taxonomic confusion due to the unusual characteristics of these dinosaurs. When initially described in 1924 Oviraptor itself was thought to be a member of the Ornithomimidae, popularly known as the "ostrich" dinosaurs, because both taxa share toothless beaks. Early caenagnathid oviraptorosaur discoveries like Caenagnathus itself were also incorrectly classified at the time, having been misidentified as birds.The hypothesis that caenagnathids were birds was questioned as early as 1956 by Romer, but not corrected until Osmolska formally reclassified them as dinosaurs in 1976. Meanwhile, the classification of Oviraptor as an ornithomimid persisted unquestioned by researchers like Romer and Steel until the early 1970s when Dale Russell argued against the idea in 1972. In 1976 when Osmolska recognized Oviraptor's relationship with the Caenagnathids, she also recognized that it was not an ornithomimid and reclassified it as a member of the former family. However, that same year Rinchen Barsbold argued that Oviraptor belonged to a distinct family he named the Oviraptoridae and he also formally named the Oviraptorosauria later in the same year.Like their classification, the paleobiology of oviraptorosaurs has been subject to controversy and reinterpretation. The first scientifically documented Oviraptor skeleton was found lying on a nest of eggs. Because its powerful parrot-like beak appeared well-adapted to crushing hard food items and the eggs were thought to belonged to the neoceratopsian Protoceratops, oviraptorosaurs were thought to be nest-raiders that preyed on the eggs of other dinosaurs. In the 1980s, Barsbold proposed that oviraptorosaurs used their beaks to crack mollusk shells as well. In 1993, Currie and colleagues hypothesized that small vertebrate prey may have also been part of the oviraptorosaur diet. Not long after, fossil embryonic remains cast doubt on the popular reconstruction of oviraptorosaurs as egg thieves when it was discovered that the "Protoceratops" eggs that Oviraptor was thought to be "stealing" actually belonged to Oviraptor itself. The discovery of additional Oviraptor preserved on top of nests in lifelike brooding posture firmly established that oviraptorosaurs had been "framed" as egg thieves and were actually caring parents incubating their own nests.